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Grubbs, S.A. 2018. An update on the stonefly fauna (Insecta, Plecoptera) of Maryland, including new and emended

state records and an updated state checklist. Illiesia, 14(04):65-80. https://doi.org/10.25031/2018/14.04

http://zoobank.org/ urn:lsid:zoobank.org:pub:D522B9EC-BAA9-49FD-AC24-

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Illiesia – http://illiesia.speciesfile.org Volume 14 – Number 4 – Page 65

AN UPDATE ON THE STONEFLY FAUNA (INSECTA, PLECOPTERA) OF

MARYLAND, INCLUDING NEW AND EMENDED STATE RECORDS AND AN

UPDATED STATE CHECKLIST

Scott A. Grubbs1

1 Department of Biology and Center for Biodiversity Studies, Western Kentucky University,

Bowling Green, Kentucky 42101, U.S.A.

E-mail: [email protected]

ABSTRACT

The stonefly fauna of Maryland is updated. An unassociated female of Perlesta Banks, 1906 is described

under informal designation, complete with light microscopy micrographs of head and pronotal

pigmentation patterns plus scanning electron microscopy (SEM) images of eggs. New state records are

presented for Cultus verticalis (Banks, 1920), Neoperla catharae Stark & Baumann, 1978, and P. mihucorum

Kondratieff & Myers, 2011. Light microscopy and SEM micrographs are also presented for P. mihucorum.

Published state records of Isoperla Banks, 1906 and Sweltsa onkos (Ricker, 1952) are emended given recent

descriptions in both genera. Distribution maps for S. onkos and S. hoffmani Kondratieff & Kirchner, 2009,

plus for all species of Isoperla and Perlesta known to occur in the Appalachian Mountain region of western

Maryland, are provided. An updated state checklist of 114 species is included.

Keywords: Plecoptera, Perlidae, Perlesta, state records, Maryland

INTRODUCTION

Information on the stonefly fauna of Maryland

was based initially on the species lists in Duffield &

Nelson (1990) and Grubbs (1997), and now easily

searchable as a “Faunal list” in DeWalt et al. (2018).

Duffield & Nelson (1990) presented 25 new state

records and the first published state checklist of 58

species. Grubbs (1997) reported 36 new state records

and an updated checklist of 95 species. Additions

and clarifications were subsequently included in

Grubbs & Stark (2001), Nelson et al. (2002), Grubbs

(2003), and Surdick (2004).

Since 2004, taxonomic and locality information

on the Maryland fauna have been included in

systematic treatments of Allocapnia Claassen, 1928

(Grubbs & Sheldon 2008), Leuctra Stephens, 1836

(Grubbs 2015, Grubbs & Wei 2017), Prostoia Ricker,

1952 (Grubbs et al. 2014), Soyedina Ricker, 1952

(Grubbs 2006), Sweltsa Ricker, 1943 (Kondratieff &

Kirchner 2009, Stark et al. 2011), Perlesta Banks, 1906

(Grubbs & DeWalt 2008, 2012), and Isoperla Banks,

1906 (Szczytko & Kondratieff 2015).

Stemming from the recent taxonomic treatments

of Sweltsa and Isoperla, several species whose

distribution in Maryland were in question, or are

now more readily identified, were re-examined.

Grubbs, S.A. 2018 An update on the stonefly fauna (Insecta, Plecoptera) of Maryland, including new and emended state records and

an updated state checklist. Illiesia, 14(04):65-80. https://doi.org/10.25031/2018/14.04


Figs. 1–6. Perlesta sp. MD-5, Maryland, Sideling Hill Creek. 1–2, female, head + pronotum, dorsal profile; 3,

entire egg, 300X; 4, egg, posterior pole, 500X; 5, egg, posterior pole and collar, 2300X; 6, egg, anterior pole

and micropyles, 600X.

Perlesta was also examined closely, revealing

unassociated females with potentially unique egg

characteristics plus a surprising range extension for

P. mihucorum Kondratieff & Myers, 2011 were

present in material collected by the author during

the 1990’s. Scanning electron and light microscopy

micrographs are included to further facilitate

identification of these two species. In addition, two

additional new state records, several species

emendations, and an updated state checklist is

provided.




METHODS

Specimens examined for this project were

obtained from Western Kentucky University,

Bowling Green (WKUC). Locality data for all

specimen records, in decimal degrees, were

obtained either directly on site with GPS units or

georeferenced from museum label data (if possible)

using Acme Mapper 2.1 (http://mapper.acme.com).

All specimen data are available as a comma separate

values file prepared in Darwin Core Archive file

format

(http://illiesia.speciesfile.org/Papers/MarylandPlec

opteraProject_DwC-A.csv). Specimens for scanning

electron microscopy (SEM) were serially

dehydrated in 75%, 95%, and 100% ethanol for 10

minutes each and placed in hexamethyldisilizane

for 30 minutes. Dehydrated specimens were

attached to aluminum stubs with double-stick tape

and coated with gold-palladium in an Emscope

SC500. Coated specimens were examined using a

Jeol JSM-6510LV scanning electron microscope and

digital images were captured with an IXRF system.

Digital images were also obtained using Auto-

Montage software with a Leica MZ16

stereomicroscope equipped with a JVC KY-F75U

digital camera.

RESULTS

Taxonomy

Perlesta MD-5

(Figs. 1–6, 21)

Female. Forewing length 9.5–11.0 mm (n = 3). Head

light pale yellow with a golden brown intraocellar

patch that is slightly open anteriorly, plus a light

golden-brown triangular patch anterior to the

anterior ocellus; m-line indistinguishable (Figs. 1–

2). Pronotum light golden brown with no medial

stripe (Figs. 1–2). Subgenital plate ca. ½ width of 8th

abdominal sternum, lacking pigmentation. Lobes

distinct, rounded laterally, bordered by a v-shaped

notch.

Egg. Oval (Fig. 3). Collar small and straight-sided,

raised about chorionic surface, flange lacking (Figs.

4–5); chorion covered by a patchwork of amoeboid-

like polygons (Figs. 4, 6); micropyles present in

anterior ¼ (Fig. 6).

Material examined. USA, Maryland, Allegany Co.,

Sideling Hill Creek, near Bellegrove, at light,

39.70443, -78.32840, 3 August 1996, S.A. Grubbs, 2♀

(WKUC); same but 14 July 1998, S.A. Grubbs, ♀

(WKUC).

Remarks. The eggs appear different from all other

species of eastern Nearctic Perlesta. A brief

description was provided here in hope that females

can be associated with males at some point in the

future. These eggs may pertain to one of three

Perlesta species whose eggs are unknown and found

along the Atlantic Coastal Plain region of Virginia

and North Carolina: P. beatyi Kondratieff, Zuellig &

Lenat, 2011, P. bjostadi Kondratieff & Kirchner, 2006,

and P. durfeei Kondratieff, Zuellig & Kirchner, 2008.

The egg of P. leathermani Kondratieff & Zuellig, 2006

also has a small, “button-like” collar (their Fig. 8.).

The light head pigmentation of MD-5 (Figs. 1–2),

however, appears different from the darker P.

leathermani (Kondratieff et al. 2006, their Fig. 1).

New state records

Neoperla catharae Stark & Baumann, 1978 is

known from several central and eastern US states

(DeWalt et al. 2018). The new records are near the

eastern end of this species range.

Perlesta mihucorum Kondratieff & Myers, 2011

was described from upstate New York and has been

collected from large streams and small rivers. This

is a common species across the panhandle region of

western Maryland plus one additional record in the

upper Piedmont region.

Remarks. Males of P. mihucorum are readily

identified by the combination of paraprocts with

subapical tooth mostly anteriorly-directed and not

visible in caudal view (Figs. 8–10) and aedeagus

with small caecum ca. as long as wide and narrow

dorsal patch (Figs. 11–12). Males from Maryland

and West Virginia examined in this study were

keyed to couplet 12 in Stark (2004), mainly because

of the first choice in couplet 10 (“Paraproct spine

distinct in lateral aspect”). This is in contrast to

Kondratieff & Myers (2011), who illustrated (their

Fig. 4) and stated “…tooth inconspicuous in lateral

view”. This initially led this author to conclude




Figs. 7–12. Perlesta mihucorum, male, Maryland, Conococheague Creek; 7, head + pronotum, dorsal profile;

8, paraprocts, lateral view, 250X; 9, paraprocts, dorsal view, 300X; 10, paraprocts, caudal view, 230X; 11,

aedeagus, dorsolateral view, 250X; 12, aedeagus, dorsal view, 250X.

that the Maryland and West Virginia specimens

represented an undescribed species. However,

paraproct tooth aspect is herein considered

interspecific variability, especially since

characteristics of the male caecum, female

subgenital plate, and egg align well with the

definition of P. mihucorum from New York.

Regarding females, this is the only species of

Perlesta with the combination of a subgenital plate

with well-developed lobes and deep v-shaped notch

(Fig. 14–15) and egg with a smooth chorion and a

wide collar that is distinctly stalked, ribbed, and

flanged apically (Figs. 16–18). Both males and

females have prominent pale yellow median bands




Figs. 13–18. Perlesta mihucorum, Maryland, Conococheague Creek; 13, female, head + pronotum, dorsal

profile; 14, female, subgenital plate, ventral view, 90X; 15, female, subgenital plate, details of left lobe,

ventral view, 400X; 16, entire egg, 270X; 17, egg, posterior pole and collar, 1200X; 18, egg, anterior pole and

micropyles, 430X.

on the pronotum (Figs. 7, 13). The eggs shown with

SEM appear inseparable as in Kondratieff & Myers

2011 (their Figs. 9–14).

The known disparate ranges of P. mihucorum as

reported from New York (Kondratieff & Myers

2011) and herein from Maryland and West Virginia

(Fig. 19) are due to incomplete understanding of the

distribution of this species. The new state records

were unexpected yet not completely surprising

considering how little is still known about Perlesta

taxonomy and biogeography. Examination of

material from the adjacent or intervening states of




Fig. 19. Distribution map of Perlesta mihucorum in eastern North America. The New York localities were

plotted using the coordinates provided in Kondratieff & Myers (2011).

Delaware, New Jersey, and Pennsylvania should

enhance understanding of the distribution of this

species. This species is also probably in northern

Virginia, more widespread in West Virginia, and

likely present in several New England states. Six

species of Perlesta in total, including the three

females conditionally determined as MD-5, are now

known from Maryland. All six species are

distributed in the small Appalachian region of

western Maryland (Figs. 19–22).

Cultus verticalis (Banks, 1920) is a widespread

Appalachian species known from Georgia

northeastward to New Hampshire and Quebec

(DeWalt et al. 2018). The females reported here were

gravid and eggs of C. verticalis are distinct from C.

decisus (Walker, 1852) (Kondratieff 2004). This state

record is not surprising.



Illiesia – http://illiesia.speciesfile.org Volume 14 – Number 4 – 1

Figs. 20–22. Distribution maps of five Perlesta species in western Maryland, USA, based on material

examined during this study. 20, P. ephelida; 21, P. nelsoni, P. placida, and P. MD sp. 5; 22, P. teaysia. MD =

Maryland, VA = Virginia, WV = West Virginia.

Species emendations and updates

Sweltsa hoffmani Kondratieff & Kirchner, 2009

and Sweltsa onkos (Ricker, 1936).

A total of 59 vials of material collected and

determined as S. onkos by the first author in the

1990s from western Maryland were reexamined.

Specimens of 55 vials, all from the Appalachian

Plateau and Ridge and Valley Physiographic

Provinces, now pertain to S. hoffmani (Fig. 23). Only

three series from the Blue Ridge Physiographic

Province (Frederick Co.) are of S. onkos (Fig. 23).

Kondratieff & Kirchner (2009, their pp. 297) likewise

listed a single series of S. onkos from Frederick

County (Fig. 23). The S. onkos collections presented

in Duffield & Nelson (1990, their Fig. 1) may also

refer to “true” S. onkos.




Fig. 23. Distribution map of Sweltsa hoffmani and S. onkos in western Maryland, USA, based on material

examined during this study plus one record in Kondratieff & Kirchner (2009). MD = Maryland, PA =

Pennsylvania, VA = Virginia, WV = West Virginia.

Isoperla Banks, 1906

The eastern Nearctic species of Isoperla were

revised by Szczytko & Kondratieff (2015), resulting

in 22 new species descriptions, revised species

definitions, and several new state records.

Consequently, all Isoperla species listed from

Maryland by Duffield & Nelson (1990) and Grubbs

(1997) require clarification and updates.

Isoperla burksi Frison, 1942 was reported from

Maryland by Duffield & Nelson (1990) (Fig. 24). The

author has not collected males of this species from

western Maryland, although undetermined females

that may refer to either I. burksi or I. orata Frison,

1942 have been found at two localities (Fig. 24).

Males with extruded aedeagi or females with

mature eggs are needed for positive determination.

Isoperla dicala Frison, 1942 was reported from

Maryland by Grubbs (2003) but not included as

occurring in the state by Szczytko & Kondratieff

(2015). The small series reported in Grubbs (2003)

from one location (Fig. 23) were determined as I.

dicala by Dr. Szczytko ca. 15 years ago.

Isoperla gibbsae Harper, 1971 was reported from

Maryland in Duffield & Nelson (1990) (Fig. 26) but

not included as occurring in the state by Szczytko &

Kondratieff (2015). This species is retained in the

Maryland state list pending verification. Szczytko &

Kondratieff (2015) similarly noted that the West

Virginia record of I. gibbsae (Kirchner 1978) requires

verification.

Isoperla holochlora Klapálek, 1923 was reported from

Maryland by Duffield & Nelson (1990) (Fig. 25).




Figs 24–25. Distribution map of four Isoperla species in western Maryland, USA, based on material examined

during this study plus records in Duffield & Nelson (1990) and Szczytko & Kondratieff (2015). 24, I. burksi

and I. montana; 25, I. dicala and I. holochlora. MD = Maryland, PA = Pennsylvania, VA = Virginia, WV = West

Virginia.




This species was included by Szczytko &

Kondratieff (2015) from the state. The author

collected this Appalachian species from western

Maryland from several localities during 1995–1997

(Fig. 25).

Isoperla kirchneri Szczytko & Kondratieff, 2015

was described from Virginia and is currently known

from New York south to North Carolina and

Tennessee (DeWalt et al. 2018). The presence of this

species in western Maryland is not surprising (Fig.

26). The specimens reported as I. namata Frison, 1942

by Grubbs (1997) were identified using Szczytko &

Kondratieff (2015) and all now refer to I. kirchneri.

Isoperla namata is herein removed from the

Maryland state list.

Isoperla montana (Banks, 1898) was reported from

Maryland by Grubbs (1997) and subsequently

listed by Szczytko & Kondratieff (2015) as

occurring in the state. The author presents several

additional localities in western Maryland (Fig. 24).

Isoperla pseudosimilis Szczytko & Kondratieff

2015 was described from upstate New York and is

currently known from the New England states

south to North Carolina and Tennessee (DeWalt et

al. 2018). This montane species was likewise expected

from Maryland and all material determined by the

author in the mid-1990s as I. similis (Hagen, 1861) were

newly determined as I. pseudosimilis using Szczytko &

Kondratieff (2015) (Fig. 27).

Isoperla similis (Hagen, 1861) was reported from

Maryland by Duffield & Nelson (1990) (Fig. 27). This

species has since been redefined by Szczytko &

Kondratieff (2015) and is now considered a species

found mainly along the Piedmont and Atlantic

Coastal Plain regions. Szczytko & Kondratieff (2015)

included I. similis from Maryland based on a single

female from the Blue Ridge Physiographic Province

(Fig. 27).

DISCUSSION

Although smaller in total land area, the updated

species total (n = 114, Table 1) reported here for

Maryland is comparable to that of Alabama (107

species, Grubbs 2011) and Georgia (100 species,

Verdone et al. 2017). Maryland is the 9th smallest

state in the US with only 21% and 24% of land area

compared to Alabama and Georgia, respectively.

Furthermore, at least 111 of the 114 species (= 97%)

have been collected from the four western

panhandle counties (Garrett, Allegany,

Washington, and Frederick) that traverse the

Appalachian Mountain region of the state. This high

proportion suggests that habitat availability and

quality is still sufficient to support a diverse

regional species pool. Elevations in the Appalachian

Plateau, Ridge and Valley, and Blue Ridge

Physiographic Provinces in western Maryland

range from ca. 60 m (= ca. 200 ft) at the Potomac

River to 1024 m (= 3360 ft) on Backbone Mountain.

The western panhandle is also sparsely populated

compared to central and eastern portions of the state

and is home to several protected areas. Notable

examples include Savage River State Forest (220

km2, Garrett Co.), Green River State Forest (192 km2,

Allegany Co.), and Catoctin Mountain Park, a

National Park Service unit in Frederick Co. (25 km2).

There are also several additional state forests, state

parks, and wildlife management areas distributed

across these four counties. By comparison, there is

markedly less surface relief in the Piedmont and

Coastal Plain Provinces within the eastern

Maryland, ranging from sea level to only 391 m (=

1282 ft) at the summit of Sugarloaf Mountain. This

region of the state is also more densely populated.

What is lacking is a broader understanding

about statewide distribution patterns of stoneflies

across Maryland. For example, how many species

with Appalachian distributions are found eastward

into the Piedmont region? How many species are

restricted in the Coastal Plain region? These

questions, and several more, can be addressed

following a comprehensive examination of material

housed in museums and private collections, a

summary of the valid literature, and focused

collection efforts in the areas of the state either

lightly sampled or supporting species clusters

indicative of important biological diversity.




Figs. 26–27. Distribution map of four Isoperla species in western Maryland, USA, based on material

examined during this study plus records in Duffield & Nelson (1990) and Szczytko & Kondratieff (2015).

26, I. gibbsae and I. kirchneri; 27, I. pseudosimilis and I. similis. MD = Maryland, PA = Pennsylvania, VA =

Virginia, WV = West Virginia.




ACKNOWLEDGEMENTS

Dr. John Andersland (Western Kentucky

University) kindly assisted with scanning electron

microscopy. Two anonymous reviewers, Bill P.

Stark, and R. Edward DeWalt provided very helpful

comments that improved this manuscript.

Table 1. List of Maryland stoneflies. New and emended records are marked by an asterisk (*)

Family Capniidae

Allocapnia aurora Ricker, 1952

Allocapnia curiosa Frison, 1942

Allocapnia frisoni Ricker & Ross, 1964

Allocapnia frumi Kirchner, 1982

Allocapnia granulata (Claassen, 1924)

Allocapnia harperi Kirchner, 1980

Allocapnia illinoensis Frison, 1935

Allocapnia maria Hanson, 1942

Allocapnia nivicola (Fitch, 1847)

Allocapnia pygmaea (Burmeister, 1839)

Allocapnia recta (Claassen, 1924)

Allocapnia rickeri Frison, 1942

Allocapnia vivipara (Claassen, 1924)

Allocapnia wrayi Ross, 1964

Allocapnia zola Ricker, 1952

Paracapnia angulata Hanson, 1961

Family Leuctridae

Subfamily Leuctrinae

Leuctra alexanderi Hanson, 1941

Leuctra carolinensis Claassen, 1923

Leuctra duplicata Claassen, 1923

Leuctra ferruginea (Walker, 1852)

Leuctra grandis Banks, 1906

Leuctra rickeri James, 1976

Leuctra sibleyi Claassen, 1923

Leuctra tenella Provancher, 1878

Leuctra tenuis (Pictet, 1841)

Leuctra variabilis Hanson, 1941

Paraleuctra sara (Claassen, 1937)

Subfamily Megaleuctrinae

Megaleuctra flinti Baumann, 1973

Family Nemouridae

Subfamily Amphinemurinae

Amphinemura delosa (Ricker, 1952)

Amphinemura nigritta (Provancher, 1876)

Amphinemura wui (Claassen, 1936)




Subfamily Nemourinae

Ostrocerca albidipennis (Walker, 1852)

Ostrocerca complexa (Claassen, 1937)

Ostrocerca truncata (Claassen, 1923)

Paranemoura perfecta (Walker, 1852)

Prostoia completa (Walker, 1852)

Prostoia similis (Hagen, 1861)

Shipsa rotunda (Claassen, 1923)

Soyedina carolinensis (Claassen, 1923)

Soyedina kondratieffi Baumann & Grubbs, 1996

Soyedina vallicularia (Wu, 1923)

Soyedina washingtoni (Claassen, 1923)

Family Taeniopterygidae

Subfamily Brachypterainae

Oemopteryx contorta (Needham & Claassen, 1925)

Strophopteryx appalachia Ricker & Ross, 1975

Strophopteryx fasciata (Burmeister, 1839)

Taenionema atlanticum Ricker & Ross, 1975

Subfamily Taeniopteryginae

Taeniopteryx burksi Ricker & Ross, 1968

Taeniopteryx lonicera Ricker & Ross, 1968

Taeniopteryx maura (Pictet, 1841)

Taeniopteryx metequi Ricker & Ross, 1968

Taeniopteryx nivalis Fitch, 1847

Taeniopteryx parvula Banks, 1918

Taeniopteryx ugola Ricker & Ross, 1968

Family Chloroperlidae

Subfamily Chloroperlinae

Alloperla aracoma Harper & Kirchner, 1978

Alloperla atlantica Baumann, 1974

Alloperla biserrata Nelson & Kondratieff, 1980

Alloperla chloris Frison, 1934

Alloperla imbecilla (Say, 1823)

Alloperla usa Ricker, 1952

Haploperla brevis (Banks, 1895)

Suwallia marginata (Banks, 1897)

*Sweltsa hoffmani Kondratieff & Kirchner, 2009

Sweltsa lateralis (Banks, 1911)

*Sweltsa onkos (Ricker, 1936)

Sweltsa palearata Surdick, 2004

Sweltsa pocahontas Kirchner & Kondratieff, 1988

Subfamily Paraperlinae

Utaperla gaspesiana Harper & Roy, 1975




Family Perlidae

Subfamily Acroneuriinae

Acroneuria abnormis (Newman, 1838)

Acroneuria arenosa (Pictet, 1841)

Acroneuria carolinensis (Banks, 1905)

Acroneuria filicis Frison, 1942

Acroneuria frisoni Stark & Brown, 1991

Acroneuria lycorias (Newman, 1839)

Attaneuria ruralis (Hagen, 1861)

Eccoptura xanthenes (Newman, 1838)

Hansonoperla appalachia Nelson, 1979

Perlesta ephelida Grubbs & DeWalt, 2012

*Perlesta mihucorum Kondratieff & Myers, 2011

Perlesta nelsoni Stark, 1989

Perlesta placida (Hagen, 1861)

Perlesta teaysia Kirchner & Kondratieff, 1997

*Perlesta sp. M-5

Perlinella drymo (Newman, 1839)

Perlinella ephyre (Newman, 1839)

Subfamily Perlinae

Agnetina annulipes (Hagen, 1861)

Agnetina capitata (Pictet, 1841)

Agnetina flavescens (Walsh, 1862)

*Neoperla catharae Stark & Baumann, 1978

Neoperla robisoni Poulton & Stewart, 1986

Neoperla stewarti Stark & Baumann, 1978

Paragnetina immarginata (Say, 1823)

Paragnetina media (Walker, 1852)

Family Perlodidae

Subfamily Isoperlinae

Clioperla clio (Newman, 1839)

Isoperla burksi Frison, 1942

Isoperla dicala Frison, 1942

Isoperla gibbsae Harper, 1971

Isoperla holochlora Klapálek, 1923

*Isoperla kirchneri Szczytko & Kondratieff, 2015

Isoperla montana (Banks, 1898)

*Isoperla pseudosimilis Szczytko & Kondratieff, 2015

*Isoperla similis (Hagen, 1861)

Subfamily Perlodinae

*Cultus verticalis (Banks, 1920)

Diploperla duplicata (Banks, 1920)

Diploperla robusta Stark & Gaufin, 1974

Isogenoides hansoni (Ricker, 1952)

Malirekus iroquois Stark & Szczytko, 1988

Remenus bilobatus (Needham & Claassen, 1925)




Yugus kirchneri Nelson, 2001

Family Peltoperlidae

Peltoperla arcuata Needham, 1905

Tallaperla elisa Stark, 1983

Tallaperla maria (Needham & Smith, 1916)

Family Pteronarcyidae

Pteronarcys biloba Newman, 1838

Pteronarcys dorsata (Say, 1823)

Pteronarcys proteus Newman, 1838

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https://doi.org/10.25031/2017/13.03

Submitted 15 February 2018, Accepted 10 April 2018, Published

30 April 2018

Hosted and published at the University of Illinois, Illinois

Natural History Survey, Champaign, Illinois, U.S.A.





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